[1] Univ Western Australia, Fac Sci, Sch Biol Sci, Oceans Inst, Crawley, WA 6009, Australia
[2] Univ Western Australia, Law Sch, Oceans Inst, Crawley, WA 6009, Australia
[3] Stanford Univ, Hopkins Marine Stn, Pacific Grove, CA 93950 USA
[4] Univ Western Australia, Fac Sci, Sch Agr & Environm, Oceans Inst, Crawley, WA 6009, Australia